Animal
Navigation
Introduction
One of the
most remarkable features of many non-human animal species is their ability
to navigate over vast distances. Examples of this animal-Olympic ability
include homing pigeons which can cover over 600 miles in one day (a feat
Virgin rail can only dream about!) and the albatross that migrates over
4000 miles. The Arctic tern accomplishes a particularly impressive feat,
although it does leave you wondering what the point is; it spends 2 weeks
at the North Pole, a few weeks at the South Pole and the rest of the year
flying between the two!
Navigation
over such vast differences may be for one of two main reasons; homing or
for the purposes of migration.
Questions on
this topic will not specify which. Typical wording would be ‘discuss
research studies into homing and/or migration in non-human species.’ This
is fortunate because the two are difficult to disentangle. Surely if an
animal migrates then on the return journey it is using similar skills and
techniques in order to get home!
However, I
shall briefly look at the reasons for homing and migration before
considering the techniques that may be used.
Homing
The texts say
little about this other than stating the obvious, i.e. that it is the
ability to find ones way home! There are clear advantages of being able
to do this. If a species has to go out looking for food or mates then it
needs to be able to find its way back to its burrow, nest etc. where
presumably it is safer. If young are involved then it is essential that
food can be taken back for them etc. The most famous species for its
homing ability has to be the ‘homing pigeon’ but others include the
salmon, purple martin and African antelope.
Migration
The reasons
for this are not always so obvious but the texts have far more to say on
the issue. Migration refers to the seasonal movement of some species
which appears to be triggered by environmental factors such as
temperature. Remember that migration is seen as a circannual rhythm.
There are
several advantages to migrating including warmer weather and avoiding
severely cold weather at the poles, new feeding grounds or watering holes,
possibility of finding new mates or sometimes avoiding predators. All of
these are clearly advantageous to a species and increase its chances of
survival or reproduction. However, migration comes at a cost. Vast
amounts of energy can be consumed and there may be many dangers, such as
predators, on the way. Fisher (1979) reported the death of at least 3,200
birds in one night in Illinois. The birds had flown into radio masts and
similar tall structures!
Birds
We normally
associate migration with bird species. It is unusual for birds to migrate
in one stage, preferring to break the journey down into smaller stages
en route. Migratory journeys by birds appear to be a combination of
innate and learned skills and this is best illustrated using the example
of Perdeck’s starlings (1958). The starlings migrate in autumn from their
breeding grounds in Russia to northern France, a south westerly journey.
Perdeck intercepted some of the starlings en route in the Netherlands and
took them south to Switzerland. Some birds were experienced others
novices on their first migration. Perdeck found that when the birds were
released to continue their journey that the young birds continued flying
in a south westerly direction which brought them out in northern Spain,
well south of where they should have been. However, the experienced birds
(no rude comments), with the benefit of past migrations behind them, were
able to adjust for the displacement and still find their way to north
France. Perdeck concluded that the young birds were relying on innate
skills whilst the mature birds were able to combine innate skills with
learning from previous experience.
Helbig (1991)
in a bizarre but ingenious experiment showed the importance of innate
factors. He took two related species of black cap, one of which migrated
south east and the other south west. When these were cross bred their
offspring, (you guessed it) flew south!
Migration in
the European stork also appears to be innate. Regardless of where the
storks originate they all migrate to the same area of north Africa.
Storks in Eastern Europe set out in an easterly direction and go via the
Middle East. Storks in Western Europe set out in a south westerly
direction and fly across the Mediterranean at Gibralter. Schuz (1971)
took eggs of the east European species and transferred them to nests in
the west. When they migrated they flew in an easterly direction just like
their biological parents.
Fish and sea
creatures
Just as birds
may use air currents such as thermals to help migrate then many sea
species use underwater currents such as the Gulf Stream to cover vast
distances. The loggerhead turtle is one such example.
Navigation
beneath the sea can be more problematic, for example it is more difficult
to use sun, moon and stars. Some research suggests that it may be
possible, for example loggerhead turtles in captivity swim towards light
sources, but generally it is assumed that their vision is not good enough
to make this a reliable method of navigation.
Salmon are
probably the fish most famed for its homing instinct. Here smell seems to
play an important role in finding natal streams (streams where they were
born). More detail about this
later.


So having
considered two reasons for why a species may need to navigate over long
distances we will now consider how they achieve this feat. This is the
most likely area to be examined!
Methods of
navigation
The simplest
method of navigation is leaving a trail that can be retraced, like
Daedulus in the Minotaur’s Labrynth (for classical scholars). The loris
(a type of lemur apparently), uses ‘urine washing’ in which they pee on
their hands and then rub the urine onto their feet to leave a scent
trail. Try it the next time you’re out!
Slightly more
complex is piloting in which landmarks are remembered en route.
These may be visual or olfactory but this method is only useful over short
distances.
Navigation by
direction
is the most complex and involves the use of sun, moon, stars and magnetic
fields to orientate yourself in relation to your destination.
In the
following section I shall only consider the last two options.
Piloting or
navigation by location
can use either visual landmarks or smells.
Landmarks
Tinbergen &
Kruyt (1938). If you’ve already revised animal memory you’ll be familiar
with this one. This is still a classic experiment and needs to be treated
as a key study. The researchers placed pine cones outside the nests of
digger wasps, a species that lives in the ground. When the wasps leave
the nest they orientate themselves by circling over the entrance to the
nest and noting the position of landmarks, in this case the strategically
placed pine cones. The researchers then move the cones a few metres away
but keeping the same pattern. On their return the wasps still try to
locate the entrance to the nest in the centre of the pine cones. As
mentioned in memory notes, the researchers ruled out the possibility of
smell being used by using a combination of scented pine cones and scented
plates.
Extension
material but good for evaluation marks (AO2):
Beusekom (1948) carried out a variation on the study placing pine cones in
a circle around the entrance to the nest. When the cones were moved they
were placed either in a circle, an ellipse or a square. They found that
the wasps would try to find the entrance in the circle and the ellipse (a
similar shape) but not in the square. In behaviourist terms the wasps had
generalised the initial learning to an ellipse (just as Little Albert
generalised from white rabbit to cotton wool etc), however they were able
to discriminate between circle and square.
Clearly
landmarks are of limited use and are only suitable for navigation over
short distances. It was once thought that racing or homing pigeons could
find their way home by following landmarks remembered when they were being
transported on their outward journey. Walcott & Schmidt-Koenig (1973)
showed that this could not be the case by anaesthetising the birds during
transportation! It is now thought that pigeons use a variety of methods
for long distance navigation and only rely on landmarks for the last bit
of the journey and locating the precise loft. (See notes on cognitive
maps in ‘animal memory’ if you require further evidence).

Cartwright &
Collett (1983) trained gerbils to find sunflower seeds and bees to find
sugar solution. They arranged the food so that it was always a fixed
distance and direction from a 40cm high cylinder. If the position of the
cylinder was moved then this confused the creatures who would search in
the wrong place suggesting:
-
That smell
was not used
-
That a
landmark (the cylinder) was being used.
However, the
two species seemed to be using different techniques. When the height of
the cylinder was altered the bees were confused. The researchers
concluded that bees were using the size of the retinal image to locate
position. However, height of the cylinder made no difference to the
gerbils’ ability to find the food. Cartwright & Collett believed that the
gerbils were using dead reckoning.
Dead reckoning
This is the
ability to know your location in respect to the target location in terms
of the distance and direction moved away from it. Even when animals have
taken a circuitous route away from the location they can still take the
shortest route back. This is like you going to Leicester via Northampton
and Coventry but coming back straight down the A6!
Olfactory maps
As already
mentioned the salmon appears to navigate its final stage of the journey
home using smell.
Much of the
research on smell has been carried out by the Italian Papi.
He believes
pigeons build up a map of their location based on smell (olfactory map).
Pigeons have
been denied their sense of smell by a variety of methods, e.g. cutting
their olfactory nerve, local anaesthetic or bunging wax up their
nostrils! This does appear to disorientate them. However, it could be
the pain and discomfort of the methods used that causes the problem.
Strangely this
disorientation only seems to occurr in Italy. It has been suggested that
pigeons here rely on smell more because they tend to be kept in lofts high
up on roof tops. In Frankfurt birds kept at ground level and deprived of
smell are able to home okay. This suggests that the way birds are
reared does affect their navigational abilities.

In more
ethical follow up studies the wind has been scented. The wind blowing
from the south is made to smell of olive oil (told you he was Italian) and
the wind blowing from the north is made to smell of turpentine (perhaps he
used to be a decorator). Pigeons then had drops of either olive oil or
turps placed on their nostrils and they flew in the direction that they
associated with that smell.
However, there
are few problems with this study, firstly it is thought pigeons have a
poor sense of smell and it is not easy to replicate due to weather
conditions.
Honey bees
also use smell to locate their own hive. Bees entering the wrong hive can
cause ‘civil unrest’ with host bees fighting off the aliens.
Fishy smells
Experiments
have been carried out on salmon returning to their natal stream. Hasler
(1986) found that plugging the nostrils of a salmon prevented it from
accurately locating its own stream. Grier & Burk (1992) exposed young
salmon to either one of two artificial smells in Lake Michigan. On their
return to the lake they entered the stream matching that smell on 90% of
occasions. It is not clear what smell the salmon are responding to under
natural conditions. It could be a case of imprinting on the
characteristic smell of that particular stream at a very young age or a
response to pheromones released by their relatives. The most likely
answer is both.
Navigation by
direction
This is the
more sophisticated method of navigation and is necessary for homing or
migrating over long distances. Possible methods available to species
include use of sun, moon and stars and magnetic fields.
Sun
Humans have
navigated using the sun for thousands of years and on clear days, even
without any complex equipment it is possible to find directions from the
position of the sun (east to west) in the sky. However, in order to do
this we also need to know the time of day. For example we know that at
midday, in the northern hemisphere, the sun is in the south and so on…
Obviously we use clocks (the clock sold by Del Boy that made him a
millionaire, was designed for navigational purposes at sea!), other
species rely on their body clocks.
Research
evidence
Bellrose
(1958) noted that on clear days (when the sun is visible) mallards take
off and immediately start heading in the right direction. However, on
overcast days they appear disorientated at the start and fly randomly
before finding their bearings.

Duck:
happy
Duck: sad
Santschi
(1911) used mirrors to reflect light from other directions and confused
the movement of ants.
Polarised
light
This is light
that has passed through a filter such as the Earth’s atmosphere.
Depending on how high in the sky the sun is more or less polarised light
gets through. When the sun is high in the sky (around midday) very little
polarisation occurs. But just after sunrise and just before sunset lots
of polarised light reaches the Earth’s surface. (See your local physicist
for more detail). It is thought that some species, for example homing
pigeons can detect polarised light and as a result can tell the position
of the sun even on days when it is obscured by cloud cover.
Von Frisch
(ultimate anorak when it comes to bees, much more on him later when we do
communication) believes that bees use polarised light to indicate position
of nectar sources in relation to the hive. It is necessary for some blue
sky to be visible for this to be possible! He confused bees by passing UV
light through a filter (creating polarisation). This caused the bees to
alter the direction of their infamous bee dances!
Clock-shifting
As I’ve
already pointed out animals rely on their biological rhythms to navigate
using the sun. These experiments are designed to alter the animals’
rhythms, confuse them into thinking it’s a different time of day, and
observing what effect this has on their navigation.
Walcott (1972)
and Keeton (1974) altered the body clocks of seagulls and pigeons
respectively. The birds are kept under artificial lighting, for example
lights come on at midnight and go off at midday, about six hours earlier
than the natural conditions outside. As a result when the birds are
released their clocks are six hours out. This equates to 90 degrees of
sun movement. As a result when the birds are transported away from their
loft and released hundreds of miles away the set off in the wrong
direction, e.g. heading north instead of east! However, they still find
their way home eventually suggesting that the sun is used as a first
resort, but that if this fails they have other methods that they can rely
on.
How birds use
the sun to navigate
Two methods
have been suggested. The map-compass hypothesis is the method already
outlined above. Animals consider the position of the sun from east to
west in the sky. So if they fly towards the sun in the evening they are
going in a westerly direction etc. The sun-arc hypothesis is more complex
because it suggests that species also consider the height of the sun in
the sky. For this to work the bird etc. must learn the position of the
sun (height and position east to west) for each time of day in its home
location. When moved away from home it is able to determine where it is
for example if the sun is lower in the sky than expected it realises that
it is further north than home etc. Grier & Burk (1992) showed that birds
only adjust for position east to west, not height of sun in the sky,
suggesting that the simpler map-compass method is used.
Stars
Bellrose
(1958) attached spotlights to the feet of mallards so he could track them
at night. He found that when the sky was clear and the stars visible that
the birds would all fly in the same direction. However, when the sky was
overcast birds would fly aimlessly. In the Northern hemisphere it appears
to be the Plough (or big Dipper for our American Cousins*) that is used as
a direction finder. The Plough is adjacent to the Pole star and rotates
around it. As a result it is always in the North.

Emlen (1975)
highlighted the importance of the plough by rearing young buntings in a
planetarium under an artificial night’s sky. In the wild the birds
migrate south in autumn and return home, in a northerly direction, in the
spring. In the planetarium the young birds appear to imprint on the
Plough and fly away from this (South) in autumn and towards it (North) in
the spring. Emlen placed ink pads and blotting paper around the bird’s
cages to record their foot prints and gauge which way they were trying to
fly.
In a follow up
experiment Emlen imprinted the birds on the star Betelgeuse (pronounced
‘beetlejuice) in the constellation of Orion. When the birds were released
into the wild they flew in the opposite direction to the one expected.
Crucially what
this does show is that although birds appear to have an innate ability to
imprint on stars for the purposes of navigation, there is still an element
of learning involved.
*Piece of
trivia: President Abraham Lincoln was watching the play ‘Our American
Cousin’ at the Ford Theatre in Washington DC when he was assassinated!
Magnetic
fields


As we have
seen birds may be temporarily disorientated by clock shifting and by
overcast skies etc., but they seem to have a back up, fail safe mechanism
for navigating if all other methods fail.
Keeton (1969)
and others have fitted magnets or Helmoltz coils (electromagnets) to the
heads of birds such as pigeons or laughing gulls and found that they
become disorientated. However, this only happened on overcast days when
the position of the sun could not be judged. Their conclusion is that
birds use the sun as their first choice but if this fails they use
magnetic fields.
Gould (1982)
reported that pigeons can become disorientated by magnetic storms and
there have been reported cases of many homing pigeons being lost when
racing during such storms.
Emlen (1976),
in an experiment similar to his planetarium study, placed young buntings
in cages in a shed. The shed had a large Helmoltz coil fitted to the
roof. Using this, Emlen was able to vary the direction of the magnetic
field inside the shed. In the spring young birds would normally jump in a
Northerly direction mimicking their migration north. However, when Emlen
adjusted the magnetic field by 120 degrees he found that the birds started
to jump in a south easterly direction instead.
How animals
detect magnetic fields
The mechanism
is not clearly understood. Beason (1989) found magnetite, a compound of
iron, in the brain of a bird called the bobolink. When magnetic fields
around the bobolink were altered using magnets, electrical activity was
recorded in these brain areas. Others however, remain sceptical.
Wiltscko & Wiltschko (1988) suggest that it may be possible for the
Earth’s magnetic fields to be detected within the visual system of some
species.
As with animal
memory, evaluation marks are tricky for this topic. Think of what the
evidence suggests and emphasise that animals seem to use different
techniques in different circumstances.
Over long
distances the sun appears to be the first choice for most species.
However, at night this is obviously not possible so the stars are used
(especially the Pole star and Plough). If conditions are overcast and sun
and stars are not visible then at least some species appear to have the
ability to use magnetic fields. Although these methods are good for
covering long distances they are not precise enough to get an animal to
its exact location.
Having got
close to their destination precise homing can be achieved using methods
such as visual landmarks or smells or both.
Other
evaluation marks can be earned by considering the possible roles played by
innate factors and learning and by criticising and/or comparing studies.
Animal
Communication
Introduction
All animals
communicate, either with members of their own species or across species.
Communication can act as a warning, a mating call or for a number of other
purposes. However, does simple communication of this type constitute
language? Hockett, and others have laid down criteria that distinguish
language from mere communication, for example a true language is able to
communicate ideas about events in the past or future, so called
displacement.
This section
looks at:
-
Natural
Animal Communication: The methods and channels various species use in
their natural environment to communicate. It considers the advantages
and disadvantages of these and looks in detail at a number of specific
examples in detail. Crucially it then considers to what extent we
believe animals possess ‘language.’
-
Human
attempts to teach language to non-human animals such as primates and
cetaceans.
1.Natural
Animal Communication
What is
communication?
Put simply it
is a two way process that allows a message to be sent and received.
Obviously for the message to be useful to both sender and receiver, the
signal sent must have the same meaning for both of them. Think of the
confusion an Englishman in New York might cause by asking for a ‘fag!’
It also seems
safe to assume that communication (or signalling) of this sort must confer
some evolutionary advantage on species as a whole, otherwise it would not
have survived as a pattern of behaviour. Individuals that use signals
would have been more likely to survive and prosper and pass their genetic
material into the next generation. However, there are examples when
signals like these can be of disadvantage to either the sender or
receiver. (see details on eaves dropping and dishonest signals).
Some possible
advantages of signals:
-
Survival.
For example warnings to other
members of the
species of an approaching
predator. (Ververt
monkey).
-
Reproduction.
Location and
attraction
of members of
the opposite sex.
(Peacock's
tail).
-
Territoriality.
Threatening gestures or
submissive
signals that settle most disputes
without
recourse to physical aggression.
(Arched back
of a domestic cat).
-
Food.
For
example letting others know
of food
locations. (Waggledance of the
honeybee).
1. Honest
signals
These usually
involve ritualised forms of normal gestures to provide a message with
unmistakeable meaning. For example the cowering of a dog to represent
submission or fear. Many mating signals also fit into this category.
Ritualisation:
Most honest signals are exaggerated forms of animal behaviour. For
example the arching of a cat or dogs back to exaggerate its height is used
as a sign of dominance to ward off would be aggressors. Similarly the
cowering posture that reduces their size is used to signal an individual’s
submissive nature. Used together these two signals can avoid costly and
aggressive encounters. Ritualised signals tend to be highly conspicuous
(and may involve lots of noise or elaborate movements). This ensures that
the signals are noticed! They also tend to be very stereotyped ensuring
that they are not misinterpreted. In some cases this can ensure that an
animal does not waste its time (and all street cred’) by attempting to
mate with the wrong species!!!
2.Dishonest
signals
These aim to
deceive and put the receiver at a disadvantage. For example smaller male
cricket frogs lower the tone of their croak to make themselves sound
larger. (A ploy used by some men in male dominated industries to make
themselves appear more macho!). The young cuckoo signalling hunger and
deceiving its adoptive parent into giving it food.
3.
Eaves-dropping
When a
predator picks up signals that are not meant for it. For example if a
signal, intended to pick up a mate, is intercepted by a predator that then
uses the information to locate the signaller. The female bark beetle (all
life is here!), releases a scent to attract males to her tree. However,
other females intercept the signal and close in on the sender and take
advantage of the attracted males she is attracting!
|
Type of signal |
Sender |
Receiver |
Example |
|
Honest Signal |
√ |
√ |
Red breast of the
vicious male robin |
|
Dishonest Signal |
√ |
x |
Hoverfly imitating
the appearance of a wasp |
|
Eaves Dropping |
x |
√ |
The bark beetle |
Channels of communication
This refers to
the sensual (broadest sense) methods that a species can use, such as
visual, auditory, tactile etc. Each has its advantages and disadvantages
and below is a list of these with specific animal examples.
Visual
It is estimated
that about 70% (some estimates put it higher) of human spoken language is
actually conveyed visually in the form of body language. In the animal
world visual messages are widely used in courtship especially in birds and
fish. The male stickleback will perform ‘zig-zag’ dance that can
stimulate a female into releasing her eggs into the water for the male to
fertilise. Robins will attack red feathers nailed to a tree but will
completely ignore a whole stuffed robin that does not possess red
colouration (Lack 1943).


Channels of
Communication
|
Method |
Advantages |
Disadvantages |
Examples |
|
Visual
Use of colour,
posture or movement.
|
Instant transmission
so good for warning of danger.
Can transmit many
messages at once, e.g. species, sex, location and level of dominance.
Directional so is
less likely to be picked up by predators.
Some, such as colour,
are permanent so are economical. |
Only useful over
short distances.
Can only be used in
daylight, unless you're a glow worm.
Can give away
precise location to predators.
Can be expensive in
terms of survival, e.g. the peacock's tail. |
Male sticklebacks
attract females by zig-zagging movements.
Male robins will
attack anything red!
Peacock's tail for
attraction of pea- hens. |
|
Olfactory
Use of pheromones.
Releasers, have a
short-term affect.
Primers, used for
long term affect such as marking territory. |
Work in the dark.
Cannot be blocked by
obstacles such as trees.
Can last for a long
time.
Not easy for
predators to understand or to interpret. |
Can be blocked by
poor weather conditions such as winds and rain.
Can take time to
produce and a relatively long time to travel to their destination.
Can last too long
and possibly get confused with later messages. |
Ants have a distinct
smell to their colony, aiding location.
Cats, mice and dogs
mark out their territories using odours.
Simmons (1990) found
that crickets avoid incest based on smell. |
|
Auditory
Species can vary
pitch, volume and sequence of sounds to transmit a variety of signals.
|
Flexible, so lots of
different messages are possible.
Sounds can travel
long distances.
Can be used in the
dark.
Sounds can be
produced quickly and travel fast (330 metres per second at sea
level!). |
Sounds can take a
lot of effort for some creatures to produce.
They are easily
detected by predators.
Distortion of sounds
occurs over long distances, especially in dense forests. |
Ververt monkey can
indicate type of predator and best response to avoid them. (see later
notes for details).
Birds tend to
communicate mainly by sound. |
|
Tactile
Touch
|
Simple to do. |
Animals have to be
close! |
Offspring of most
bird species request food this way.
Grooming during
courtship.
|
Auditory (sounds)
As humans
(speaking for the majority of us now), this is the channel we most
associate with language and communication. Many other species, most
notably birds, also make good use of sounds in communication and
signalling.
Sounds can be
varied in a number of ways:
·
Pitch (or tone): female
toads apparently prefer males with a deep ‘voice’ as they suggest larger
males (obviously no one has told them that size doesn’t matter Ed).
·
Volume: clearly a louder
signal will have more impact and travel further.
·
Sequence: the order in
which the sounds are deployed, crucial in human language but also in other
species such as crickets.
Many species
vary all three to good effect to alter the meaning of their call. A good
example of this is the vervet monkey that we will look at in more detail
later.
 |
Great
tits (an example of deceitful use of auditory signals)
Males of
the species change their song each time they change perch (move from
one branch or tree to the next).
Krebs (1977) believed that they used this to con other males
into believing that there were already lots of males in the area and
that breeding opportunities would be limited.
However, Yasukawa (1981), believed that other males may realise
that there is only one male in the area, however, due to his
repertoire he must be strong and experienced so more than a match for
them! |
Olfactory
(smells)
Pheromones
(chemical messengers) are usually the method of first choice!
Releaser
pheromones usually have a short term effect bringing about a sudden change
in behaviour, for example attracting male moths to a female releaser.
Primer
pheromones usually have a longer term effect and may alter the physiology
of the receiver. It is common for many species such as domestic cats to
mark their territory with scent. This is achieved by the pheromones in
their urine.
A few
statistics: each antenna of the male silk worm moth has 10,000 hairs that
it uses to detect female pheromones. Just a few molecules can change the
behaviour of the moth. Simmons (1990) found that smell can be crucial in
preventing accidental incestuous breeding. Female crickets showed a
preference for more unrelated males as evidenced by their pheromone.
Attachments
between mother and offspring may also be mediated by smell. Farmers will
cover an orphaned lamb in the afterbirth of another newly born lamb to
persuade the mother to adopt the orphan. (Once saw this being done… then
went home and had a neck of lamb casserole that I’d made earlier!)
What is
language?
Psychologists
as well as linguists have problems in defining ‘language,’ both finding it
easier and more useful to identify the different properties that
characterise language. The most widely used set of criteria are those
devised by linguist Charles Hockett who has compared human languages with
other forms of communication.
Hockett’s
criteria:
Symbolic or
semanticity: the method of communication uses symbols that have a
shared meaning between all those members of the species using it. In
human terms, the word ‘tree’ in English, has a shared meaning between all
people around the world that speak our language.
Syntax:
the use of these symbols requires rules, for example in English the
adjective usually goes before the noun ‘the red book’ as opposed to ‘the
book red.’ Those that do French will be aware that this is not always the
case in French. Some adjectives are placed after the noun.
Arbitrariness: the symbols used bear no resemblance to the action or
object that they are representing. The word ‘car’ is arbitrary since it
looks or sounds nothing like the object that it represents. The
waggledance of the bee however is not arbitrary since the direction of the
dance represents the direction of the nectar and the speed of the dance
reflects the distance.
Specialisation: the sounds created have no other function other than
what they are representing. For example the panting of a dog has a
biological purpose. A dog squealing because of pain does not do so to
communicate the pain but because it is in pain.
Displacement:
the language can communicate about actions, objects or emotions that are
not present or visible at that moment. For example the waggledance shows
displacement because it refers to nectar not visible to the dancer. Human
languages can communicate ideas about actions that occurred yesterday or
may happen tomorrow, so are not impinging on the individual at that
moment. Most animal communication refers to immediate environmental
stimuli such as the presence of a predator.
Cultural
transmission: the method of communication is passed from one
generation to the next by a process of teaching. This appears to be the
case with some birdsong but is not true of the waggledance which is innate
and present from birth.
Generativity
or productivity: the number of utterances possible using the language
is infinite. Using the English language I could say ‘Me and Kylie popped
down the Sugarloaf for a pint of Abbot and a prawn vindaloo.’ The chances
are nobody has ever said that before. Most methods of animal
communication have nothing like that level of flexibility. The calls of
most species are very limited in scope.
Prevarication: the language can be used to tell lies or jokes.
Discreteness: the language combines smaller units (e.g. words) to
create meaning (e.g. sentences).
Interchangeability: an individual can both send and receive messages.
Hockett’s
first criterion, not mentioned above, is that the language should be
vocal or auditory. I leave this ‘til last since it not only rules out
the waggledance, but would appear to relegate nearly all attempts of
teaching apes and cetaceans to mere communication. It would also rule out
sign language!!!
Does natural
animal communication constitute language?
What follows is
a brief description of various natural signalling systems and a
consideration of whether or not they fulfil Hockett's criteria.
1. Birds
Birds make most
use of the auditory channel, so called birdsong. This is often used in
conjunction with other channels such as visual signalling. Hunter and
Krebs (1979) found that the nature of their song relates to their
environment.
·
In open spaces birds use
a wider range of frequencies and repeat notes and sequences of sound
faster.
·
In dense forests they
use lower frequencies.
Wiley &
Richards (1978) attributed this to communication of the message with
minimum distortion. In forests trees cause reverberations. Lower pitched
sounds are less likely to be disrupted. In open spaces the greatest risk
is from strong winds. High-pitched sounds, quickly repeated are less
likely to be affected.
Is birdsong
innate or learned? (Easy evaluation marks to be had here).
Crickets reared
in isolation (so they have never heard other crickets sing), still sing
themselves. Obviously, crickets are not birds, but this suggests that
their song is innate. However, higher species, such as sparrows, when
reared in isolation between 8 and 90 days old, fail to pick up birdsong,
suggesting that it is learned, or as it applies to Hockett, ‘culturally
transmitted.’
Note. It is
possible for birds reared like this to pick up the song of related
species. The conclusion, therefore is, that the ability to sing is
innate, the nature of their song is learned.
|
Criterion |
Yes or no |
Comments |
|
Symbolic |
Yes |
Some sounds
represent anger, others courtship etc. |
|
Specialised |
Yes |
These are different
to sounds made in pain etc. |
|
Generative |
No |
Very limited number
of meanings to sounds. |
|
Syntax |
No |
Too few sounds used
to require rules |
|
Critical period |
Yes |
Birds need to be
exposed to language at a young age |
|
Cultural
transmission |
Yes |
Signals passed onto
young. |
Is it
language? No.
2. Honeybee
The dances of
the honeybee were studied by von Frisch over a period of many years. Two
distinct types of dance were observed:
1. The Round
dance. (Indicating nectar within an 80m radius).
The returning
bee dances in a circle, as the name suggests. The other bees then fly off
and search nearby. This dance gives no indication of direction.
2. The
Waggledance. (Indicating nectar more than 80m away).
The returning
bee performs a more elaborate dance that indicates approximate distance,
and crucially direction.
a. Direction
is indicated by the angle at which the dance is performed. The dance
comprises of a figure of eight. The straight stretch in the middle is the
relevant bit. If this is vertical on the wall of the hive it informs the
others that the nectar is towards the sun. Dancing downwards would mean
fly away from the sun etc.
b.
Distance is indicated by
the energy put into the dance:
i.
Number of times the bee
completes the cycle
ii.
Number of waggles
iii.
Amount of noise made.
The greater the energy expenditure the
nearer the nectar is to the hive. Remember that the hive is dark inside
so visibility is minimal. The observing bees therefore follow the dancer
to assess direction and the dancer herself regurgitates some of the nectar
as an additional clue.
Subsequent
research has backed up von Frisch’s early work on the complex nature of
the dance. The method of communication has some degree of flexibility.
For example the bees only dance on about 10% of occasions when the source
they have found is particularly plentiful or if the find satisfies a
particular need of the hive.
The receivers
don’t always act on the information. The Goulds sat in a boat in the
middle of a lake and provided nectar to passing bees. These returned to
the hive and performed the appropriate dance communicating the location of
the find. However, the others did not act upon the information. The
Goulds assumed this was due to the bees having a mental or cognitive map
of their immediate environment. They would have realised that the dance
was indicating the presence of nectar in the middle of water. Since this
would normally be impossible the receivers assume a mistake has been made
and ignore the message.
|
Criterion |
Yes or no? |
Comments |
|
Symbolic |
Yes |
Uses angle and
vigour to represent direction and distance. |
|
Specialised |
Yes |
The dance has no
other uses. |
|
Displacement |
Yes |
The nectar is out of
sight. |
|
Syntax |
No |
Not needed. |
|
Cultural
transmission |
No |
The dance is innate
and does not develop over time. |
|
Arbitrariness |
No |
Vigour of dance is
proportional to distance from hive. |
Is it
language? No.
Additional
points
Bee dances are
not productive in that the message is always communicating the same thing,
no new subjects are incorporated. Also the language does not demonstrate
reflexive in that the bees are unable to communicate anything about
themselves.
3.
Whales
Whales
communicate via song and this is often compared to the songs of birds.
Typically a song lasts about 30 minutes and comprises long, slow notes.
Songs are split into themes and themes into phrases. Finally each phrase
comprises notes. Whale species average about six themes, but they do
change over time. All the whales in a given area sing the same song but
this does change during the course of a season. At the start of the next
singing season the whales sing the same song as they were singing at the
end of the previous season. The meaning of the songs is difficult to
interpret and a number of suggestions have been put forward. Some have
suggested that given the huge brain of the whale its songs must have
complex meanings, but this appears not to be the case.
1. Mating
call. Winn & Winn (1985), along with others, have reported that only
males sing suggesting a mating role for the songs, seeking to attract
females. They suggest that a build up of androgen (male hormone) triggers
the call. Tyack (1981) watched singers pursue non singers and then engage
in courtship type behaviour, again suggesting a mating role.
2. Warding
off other males. Winn & Winn (1985) suggest that the lower frequency
notes of the songs may be an attempt by males to keep other males at bay.
Typically songs combine notes of different pitch, so the songs could be
conveying different messages.
3. Feeding
behaviour. (D’Vincent 1985) suggest songs appear to play a vital role in
all manner of social behaviours including feeding.
4. Surfacing.
Whales need to surface at regular intervals, Winn et al (1979) report a
‘ratcheting sound’ immediately prior to surfacing and this has enabled
scientists to predict when whales will surface.
It is worth
remembering that the song of the humpback whale will save the earth in the
23rd century! (Information published courtesy of the producers
of Star Trek).
|
Criterion |
Yes or no? |
Comments |
|
Symbolic |
| |